How honeybees make grazing landings on flat surfaces

 Freely flying bees were filmed as they landed on a flat, horizontal surface, to investigate the underlying visuomotor control strategies. The results reveal that (1) landing bees approach the surface at a relatively shallow descent angle; (2) they tend to hold the angular velocity of the image of the surface constant as they approach it; and (3) the instantaneous speed of descent is proportional to the instantaneous forward speed. These characteristics reflect a surprisingly simple and effective strategy for achieving a smooth landing, by which the forward and descent speeds are automatically reduced as the surface is approached and are both close to zero at touchdown. No explicit knowledge of flight speed or height above the ground is necessary. A model of the control scheme is developed and its predictions are verified. It is also shown that, during landing, the bee decelerates continuously and in such a way as to keep the projected time to touchdown constant as the surface is approached. The feasibility of this landing strategy is demonstrated by implementation in a robotic gantry equipped with vision. Received: 21 June 1999 / Accepted in revised form: 20 March 2000     

Optic flow cues guide flight in birds

Although considerable effort has been devoted to investigating how birds migrate over large distances, surprisingly little is known about how they tackle so successfully the moment-to-moment challenges of rapid flight through cluttered environments [1]. It has been suggested that birds detect and avoid obstacles [2] and control landing maneuvers [3-5] by using cues derived from the image motion that is generated in the eyes during flight. Here we investigate the ability of budgerigars to fly through narrow passages in a collision-free manner, by filming their trajectories during flight in a corridor where the walls are decorated with various visual patterns. The results demonstrate, unequivocally and for the first time, that birds negotiate narrow gaps safely by balancing the speeds of image motion that are experienced by the two eyes and that the speed of flight is regulated by monitoring the speed of image motion that is experienced by the two eyes. These findings have close parallels with those previously reported for flying insects [6-13], suggesting that some principles of visual guidance may be shared by all diurnal, flying animals.     

Effects of seed morphology and orientation on secondary seed dispersal by wind

传播体形态与方位对二次种子风力传播的影响     

Behaviour of coconut mites preceding take-off to passive aerial dispersal

For more than three decades the coconut mite Aceria guerreronis Keifer is one of the most important pests of coconut palms and has recently spread to many coconut production areas worldwide. Colonization of coconut palms is thought to arise from mites dispersing aerially after take-off from other plants within the same plantation or other plantations. The underlying dispersal behaviour of the mite at take-off, in the airborne state and after landing is largely unknown and this is essential to understand how they spread from tree to tree. In this article we studied whether take-off to aerial dispersal of coconut mites is preceded by characteristic behaviour, whether there is a correlation between the body position preceding aerial dispersal and the direction of the wind, and whether the substrate (outer surface of coconut bracts or epidermis) and the wind speed matter to the decision to take-off. We found that take-off can sometimes be preceded by a raised body stance, but more frequently take-off occurs while the mite is walking or resting on its substrate. Coconut mites that become airborne assumed a body stance that had no relation to the wind direction. Take-off was suppressed on a substrate providing food to coconut mites, but occurred significantly more frequently on the outer surface of coconut bracts than on the surface of the fruit. For both substrates, take-off frequency increased with wind speed. We conclude that coconut mites have at least some degree of control over take-off for aerial dispersal and that there is as yet no reason to infer that a raised body stance is necessary to become airborne.     

Factors influencing the landing of maleEpiphyas postvittana (Walker) exhibiting pheromone-mediated flight (Lepidoptera: Tortricidae)

The effects of changes in various visual and olfactory properties of a white card surface on the landing position of male Epiphyas postvittanaexhibiting pheromone-mediated flight were studied in a wind tunnel. Males landed predominantly at the most downwind position of a surface in line with the pheromone source, regardless of the strength of the source. The position on the surface that males landed was strongly influenced by visual factors. The landing position of males appeared to be influenced by visual cues along all three axes of the surface. Decreases in either the dimension horizontally perpendicular to the wind direction or the vertical dimension resulted in greater numbers of males landing farther upwind on the surface than the downwind edge. Visual changes in the axis along the wind direction also affected the position at which males landed. For example, when presented with two white card surfaces with a 4- cm gap between them, males tended to land on the downwind edge of the upwind surface (on which the source was located). When the gap was bridged with clear Mylar, the landing pattern was significantly different, with the greater proportion of males landing on the downwind surface. However, when Mylar was placed on the plexiglass floor of the tunnel (in addition to bridging the gap), the landing pattern on the surface was not significantly different from that on the two surfaces without the Mylar bridge. It is suggested that during the prelanding and landing phases of pheromone-mediated flight, male moths orient to visual features of the surface containing the pheromone source rather than to visual features of the source (conspecific female moth) itself.     

A bio-inspired flying robot sheds light on insect piloting abilities

When insects are flying forward, the image of the ground sweeps backward across their ventral viewfield and forms an "optic flow," which depends on both the groundspeed and the groundheight. To explain how these animals manage to avoid the ground by using this visual motion cue, we suggest that insect navigation hinges on a visual-feedback loop we have called the optic-flow regulator, which controls the vertical lift. To test this idea, we built a micro-helicopter equipped with an optic-flow regulator and a bio-inspired optic-flow sensor. This fly-by-sight micro-robot can perform exacting tasks such as take-off, level flight, and landing. Our control scheme accounts for many hitherto unexplained findings published during the last 70 years on insects' visually guided performances; for example, it accounts for the fact that honeybees descend in a headwind, land with a constant slope, and drown when travelling over mirror-smooth water. Our control scheme explains how insects manage to fly safely without any of the instruments used onboard aircraft to measure the groundheight, groundspeed, and descent speed. An optic-flow regulator is quite simple in terms of its neural implementation and just as appropriate for insects as it would be for aircraft.     

Reducing ground reaction forces in gymnastics’ landings may increase internal loading

The aim of this study was to use a subject-specific seven-link wobbling mass model of a gymnast, and a multi-layer model of a landing mat, to determine landing strategies that minimise ground reaction forces (GRF) and internal forces. Subject-specific strength parameters were determined that defined the maximum voluntary torque/angle/angular velocity relationship at each joint. These relationships were used to produce subject-specific ‘lumped’ linear muscle models for each joint. Muscle activation histories were optimised using a Simplex algorithm to minimise GRF or bone bending moments for forward and backward rotating vault landings. Optimising the landing strategy to minimise each of the GRF reduced the peak vertical and horizontal GRF by 9% for the backward rotating vault and by 8% and 48% for the forward rotating vault, compared to a matching simulation. However, most internal loading measures (bone bending moments, joint reaction forces and muscle forces) increased compared to the matching simulation. Optimising the landing strategy to minimise the peak bone bending moments resulted in reduced internal loading measures, and in most cases reduced GRF. Bone bending moments were reduced by 27% during the forward rotating vault and by 2% during the backward rotating vault landings when compared to the matching simulations. It is possible for a gymnast to modify their landing strategy in order to minimise internal forces and lower GRF. However, using a reduction in GRF, due to a change in landing strategy, as a basis for a reduction in injury potential in vaulting movements may not be appropriate since internal loading can increase.     

The influence of tactile and other non-chemical factors on the ovipositional responses of the generalist herbivore Epiphyas postvittana

A bioassay to test the ovipositional responses of the generalist herbivore, Epiphyas postvittana, was developed, and various non-chemical factors that influenced the laying, and number of eggs laid, by females were tested. In choice experiments using various models, surface texture was found to be an important factor influencing ovipositional behaviour, with females preferring to lay on smoother rather than rougher surfaces. Another surface texture feature that influenced oviposition was a raised wax ridge on a smooth surface. The wax ridge stimulated more eggs to be laid within close proximity to the ridge. Observations of females showed that the increased numbers of eggs laid on models with a wax ridge resulted not from increased landing, time spent, or numbers of eggs laid per ovipositional bout, but rather through an increased probability of ovipositing after landing on models with a wax ridge compared to models without the ridge. Spatial features of the models were also found to influence egglaying by females. Females showed a tendency to lay on the upper rather than the lower surface of models, although this appeared to depend upon how stimulatory the upper surface of the model was. Females also laid more eggs on models with a dense array of leaves than on models with a more spread-out array.     

The daily pattern of autumn bird migration in the northern Sahara

The temporal pattern of migration by passerine birds during the night, and their arrival during the day at the Egyptian coast and in the northern Sahara Desert was investigated. The mean direction of nocturnal migration at the coast was south-southeast, while at all desert sites it was south-southwest.
Birds arrived at the Egyptian coast only during the second half of the night which is explained by the fact that no birds could have taken off from the Mediterranean Sea. At least some of the birds landed at the coast where they spent the day before taking off shortly after sunset. These birds passed the desert sites at the expected time of day assuming a ground speed of 18 m per second. However, the origin of the birds passing the desert sites early at night is unclear. They must either have spent the day in the desert north of the study sites or they had overflown the Egyptian coast in the afternoon without landing.
The landing of birds during the day at the desert sites was bimodal. This pattern of arrival is explained either by some birds having landed at the Egyptian coast in the early morning before continuing, or by deteriorating conditions later in the day during flight or when resting in the desert, that obliged them to seek shelter at the desert sites.
A correlation between the number of migrants observed during the night and the number of resting birds in the desert on the following day suggests that an unknown proportion of birds might regularly use an intermittent migratory strategy that includes rest periods by day when crossing the desert, whereas others might adapt a non-stop migratory strategy.     

Three-dimensional analysis of aphid landing behaviour in the laboratory and field

The final second of the landing approach of black bean aphids, Aphis fabae, was analysed in three dimensions using video techniques. A yellow landing platform was placed upwind or downwind from aphids aggregating under a ceiling light in a laboratory wind tunnel with 10, 20, 30, 40 or 50 cm s^–1 wind speeds, and up-tunnel or down-tunnel in still air. As individual aphids flew to the platform, body orientation (assessed by direct observation) was predominantly into-wind whether the initial flight direction to the landing platform was upwind or downwind. A greater proportion showed into-wind body orientation as wind speed increased. Flight track parameters which differed significantly between wind speeds were the track length, linear start to finish distance, linearity index, horizontal ground speed, speed vertical to the ground, vertical turning rate, and horizontal turning rate. The position of the landing platform was important for track length, linear start to finish distance, horizontal ground speed, three-dimensional turning rate, horizontal turning rate, vertical turning rate, and sinuosity. As wind speed increased above 30 cm s^–1 the ground speed became more consistent and indicated considerable variation in air speed to adjust for ground speed. For the majority of aphids there was a strong preference (88%) for into-wind landings with initial upwind directed flight, while for downwind flights a significant number (55%) of insects reversed initial flight direction and landed into-wind. Field recorded landings showed that 66% of aphids landed into-wind and there was a mean bearing to the wind of 71 ± 42°, a similar finding to wind-tunnel studies.     

Object detection by honeybees: Why do they land on edges?

Behavioural experiments using a variety of experimental situations (Figs. 1, 3, 6, 7, 9) were conducted to investigate the visual cues which bees use in the task of object-ground discrimination. The bees' flight and landing behaviour was video-filmed throughout the experiments. The evaluation of the video data shows that bees trained to find a randomly textured figure raised above a similarly textured ground land mainly on the boundaries of the figure, facing its inner surface (Fig. 2a, b). Bees can also be trained to find a hole, i.e. a low texture viewed through a window cut in a raised texture, but these bees are not attracted to the edges of the hole (Fig. 5a, b). Bees trained to a single edge between a low and a raised random texture land at the edge mainly facing the raised side (Table 1). Bees approaching the edge from the high side cross the edge in most cases without landing on it (Table 1). Bees trained to an edge between 2 striped patterns, one raised above the other, again land on the edge facing the raised pattern, regardless of whether the stripes on the 2 patterns run parallel or perpendicular to each other or to the edge (Fig. 8). In this case, the bees acquire range information by flying in oblique directions with respect to the orientation of the stripes (Fig. 10). All of the results suggest that the edge elicits landings when the bee perceives a local increase in the speed of image motion, signalling an abrupt decrease in range. This is corroborated by the results of further experiments in which artificial motion was used to simulate range differences between the two sides of an edge (Table 2). We conclude that image speed is a powerful cue in range discrimination as well as object detection. Dedicated to G. Adrian Horridge on the occasion of his retirement     

A multi-differential neuromorphic approach to motion detection

This paper presents a multi-differential neuromorphic approach to motion detection. The model is based evidence for a differential operators interpretation of the properties of the cortical motion pathway. We discuss how this strategy, which provides a robust measure of speed for a range of types of image motion using a single computational mechanism, forms a useful framework in which to develop future neuromorphic motion systems. We also discuss both our approaches to developing computational motion models, and constraints in the design strategy for transferring motion models to other domains of early visual processing.     

Elevated gastrocnemius forces compensate for decreased hamstrings forces during the weight-acceptance phase of single-leg jump landing: implications for anterior cruciate ligament injury risk

Approximately 320,000 anterior cruciate ligament (ACL) injuries in the United States each year are non-contact injuries, with many occurring during a single-leg jump landing. To reduce ACL injury risk, one option is to improve muscle strength and/or the activation of muscles crossing the knee under elevated external loading. This study?s purpose was to characterize the relative force production of the muscles supporting the knee during the weight-acceptance (WA) phase of single-leg jump landing and investigate the gastrocnemii forces compared to the hamstrings forces. Amateur male Western Australian Rules Football players completed a single-leg jump landing protocol and six participants were randomly chosen for further modeling and simulation. A three-dimensional, 14-segment, 37 degree-of-freedom, 92 muscle-tendon actuated model was created for each participant in OpenSim. Computed muscle control was used to generate 12 muscle-driven simulations, 2 trials per participant, of the WA phase of single-leg jump landing. A one-way ANOVA and Tukey post-hoc analysis showed both the quadriceps and gastrocnemii muscle force estimates were significantly greater than the hamstrings (p<0.001). Elevated gastrocnemii forces corresponded with increased joint compression and lower ACL forces. The elevated quadriceps and gastrocnemii forces during landing may represent a generalized muscle strategy to increase knee joint stiffness, protecting the knee and ACL from external knee loading and injury risk. These results contribute to our understanding of how muscle?s function during single-leg jump landing and should serve as the foundation for novel muscle-targeted training intervention programs aimed to reduce ACL injuries in sport.     

Surprising absence of association between flower surface microstructure and pollination system

• The epidermal cells of flowers come in different shapes and have different functions, but how they evolved remains largely unknown. Floral micro‐texture can provide tactile cues to insects, and increases in surface roughness by means of conical (papillose) epidermal cells may facilitate flower handling by landing insect pollinators. Whether flower microstructure correlates with pollination system remains unknown.
• Here, we investigate the floral epidermal microstructure in 29 (congeneric) species pairs with contrasting pollination system. We test whether flowers pollinated by bees and/or flies feature more structured, rougher surfaces than flowers pollinated by non‐landing moths or birds and flowers that self‐pollinate.
• In contrast with earlier studies, we find no correlation between epidermal microstructure and pollination system. The shape, cell height and roughness of floral epidermal cells varies among species, but is not correlated with pollinators at large. Intriguingly, however, we find that the upper (adaxial) flower surface that surrounds the reproductive organs and often constitutes the floral display is markedly more structured than the lower (abaxial) surface.
• We thus conclude that conical epidermal cells probably play a role in plant reproduction other than providing grip or tactile cues, such as increasing hydrophobicity or enhancing the visual signal.

     

Unifying shading and texture through an active observer

Shading (variations of image intensity) provides an important cue for understanding the shape of three-dimensional surfaces from monocular views. On the other hand, texture (distribution of discontinuities on the surface) is a strong cue for recovering surface orientation by using monocular images. But given the image of an object or scene, what technique should we use to recover the shape of what is image? Resolution of shape from shading requires knowledge of the reflectance of the imaged surface and, usually, the fact that it is smooth (i.e. it shows no discontinuities). Determination of shape from texture requires knowledge of the distribution of surface markings (i.e. discontinuities). One might expect that one method would work when the other does not. I present a theory on how an active observer can determine shape from the image of an object or scene regardless of whether the image is shaded, textured, or both, and without any knowledge of reflectance maps or the distribution of surface markings. The approach is successful because the active observer is able to manipulate the constraints behind the perceptual phenomenon at hand and thus derive a simple solution. Several experimental results are presented with real and synthetic images.     

Classification and comparison of biomechanical response strategies for accommodating landing impact

The purposes of this study were to (a) present a theoretical model to explain the methods by which individuals accommodate impact force in response to increases in an applied stressor, (b) use the model and a correlation procedure to classify a sample of individuals based on their observed response patterns, and (c) statistically evaluate the classification process. Ten participants performed landings from three heights while video and force platform data were being collected. Magnitudes of impact-force characteristics from ground reaction force and lower extremity joint moments were evaluated relative to changes in landing momentum. Correlation between impact force and landing momentum was used to classify participant responses into either a positive or negative biomechanical strategy, as defined by the model. Positive and negative groups were compared using the Mann-Whitney U test. Results indicated that all responses fit within the categories defined by the model. Some individuals preferred positive strategies while others preferred negative ones depending on the specific variable. Only one participant consistently exhibited the negative strategy for all variables. Positive and negative groups were determined to be statistically different, p < or = 0.05, for 61% of the comparisons, suggesting actual differences between groups. The proposed model appeared robust and accounted for all responses in the current experiment. The model should be evaluated further using landing and other impact activities; it should be refined and used to help researchers understand individual impact-response strategies in order to identify those who may be at risk for impact related injuries.     

Anuran metamorphosis: a model for gravitational study on motor development

Limbs and supporting structures of an organism experience a full weight of its own when it lands from water, because neutral buoyancy in the aquatic habitat will be no longer available in the terrestrial world. Metamorphosis of anuran amphibians presents a good research model to examine how this transition from non-loading to weight-loading affects development of motor capacity at the time of their first emergence on land. Our video analysis of the transitional anurans, Rana catesbeiana, at Gosner stage 46 (the stage of complete transformation) demonstrated that the take-off speed increased 1.23-fold after the first six hours of weight-loading on the wet ground. It did not increase further during the following three days of loading, and was close to the level of mature frogs with different body mass. During development of larvae in deep water with no chance of landing through metamorphosis, both tension and power of a hindlimb anti-gravity muscle increased 5-fold between stages 37 and 46. However, the muscle contractility increased more rapidly when the larvae could access the wet ground by their natural landing behavior after stages 41-42. Muscle power, one of major factors affecting locomotory speed, was 1.29-fold greater in the loaded than in the non-loaded larvae at the transitional stage. Thus, weight-loading had a potentially significant effect on the elevation of motor capacity, with a similar extent of increment in locomotory speed and muscle power during the last stages of metamorphosis. Such a motor adjustment of the froglets in a relatively short transitional period would be important for effective ecological interactions and survival in their inexperienced terrestrial life.     

Passive dynamics change leg mechanics for an unexpected surface during human hopping.

Humans running and hopping maintain similar center-of-mass motions, despite large changes in surface stiffness and damping. The goal of this study was to determine the contributions of anticipation and reaction when human hoppers encounter surprise, expected, and random changes from a soft elastic surface (27 kN/m) to a hard surface (411 kN/m). Subjects encountered the expected hard surface on every fourth hop and the random hard surface on an average of 25% of the hops in a trial. When hoppers on a soft surface were surprised by a hard surface, the ankle and knee joints were forced into greater flexion by passive interaction with the hard surface. Within 52 ms after subjects landed on the surprise hard surface, joint flexion increased, and the legs became less stiff than on the soft surface. These mechanical changes occurred before electromyography (EMG) first changed 68-188 ms after landing. Due to the fast mechanical reaction to the surprise hard surface, center-of-mass displacement and average leg stiffness were the same as on expected and random hard surfaces. This similarity is striking because subjects anticipated the expected and random hard surfaces by landing with their knees more flexed. Subjects also anticipated the expected hard surface by increasing the level of EMG by 24-76% during the 50 ms before landing. These results show that passive mechanisms alter leg stiffness for unexpected surface changes before muscle EMG changes and may be critical for adjustments to variable terrain encountered during locomotion in the natural world.     

An experimental and analytical study of impact forces during human jumping

Impact forces during landing in dismounts from the horizontal bar onto regulation gymnastic mats and in jumping from a height of 0.45 m onto a hard surface were measured. A two degree-of-freedom dynamic model was developed to predict the forces in landing on the hard surface. The periods of the two peaks that can be identified from experimental data were used in the determination of the system parameters. The peak forces recorded in gymnasts' landing ranged from 8.2 to 11.6 times the body weight. Maximum forces in jumping from 0.45 m, which ranged from 5.0 to 7.0 times the body weight, were accurately predicted by the model.     

Decreased non-MHC-restricted (CD56+) killer cell cytotoxicity after spaceflight.

Cytotoxic activity of non-major histocompatibility complex-restricted (CD56+) (NMHC) killer cells and cell surface marker expression of peripheral blood mononuclear cells were determined before and after spaceflight. Ten astronauts (9 men, 1 woman) from two space shuttle missions (9- and 10-day duration) participated in the study. Blood samples were collected 10 days before launch, within 3 h after landing, and 3 days after landing. All peripheral blood mononuclear cell preparations were cryopreserved and analyzed simultaneously in a 4-h cytotoxicity (51)Cr release assay using K562 target cells. NMHC killer cell lytic activity was normalized per 1,000 CD56+ cells. When all 10 subjects were considered as one study group, NMHC killer cell numbers did not change significantly during the three sampling periods, but at landing lytic activity had decreased by approximately 40% (P < 0.05) from preflight values. Nine of ten astronauts had decreased lytic activity immediately after flight. NMHC killer cell cytotoxicity of only three astronauts returned toward preflight values by 3 days after landing. Consistent with decreased NMHC killer cell cytotoxicity, urinary cortisol significantly increased after landing compared with preflight levels. Plasma cortisol and ACTH levels at landing were not significantly different from preflight values. No correlation of changes in NMHC killer cell function or hormone levels with factors such as age, gender, mission, or spaceflight experience was found. After landing, expression of the major lymphocyte surface markers (CD3, CD4, CD8, CD14, CD16, CD56), as determined by flow cytometric analysis, did not show any consistent changes from measurements made before flight.